Water Regimes and Ecological Adaptations 63 \_-2rTand or terrestrial plants. They grow in. normal water conditions with their assimilatory organs adapted to existence in air, Land plants exhibit many grades of adaptation to their mode of life, Thus, those which encounter greatest difficulties in securing water are the xerophytes, while others as mesophytes, which in some respects occupy an intermediate position between the two extremes, the hydyophytes and the xerophytes. ‘3. Helophytes or marshy plants. They are an intermediate group between acquatic and land plants. An account of these ecological groups of plants, i.e, hydrophytes, mesophytes, xerophytes and halophytes will be presented here, ‘They grow on extremely wet soil where water is available to plants in abundance. According to the way in which they develop in water, they are further subdivided into the following five Categories: 1. Free-Floating Hydrophytes They remain in contact with water and air, but Rot soil. They float freely on the water surface, Leaves in some are very minute, while in others quite large. Some of the free-floating hydrophytes, as Wolffia, Lemna, Spirodella, Azolla, Kichhornia, Salvinia and Pistia are shown in Figure 4.2. 2. ‘Rooted Hydrophytes with Floating Leaves Their roots are fixed in mud, but leaves have long petioles which keep them floating on the water surface, The remaining plant, except leaves, remains in water. Some of the rooted hydrophytes with’ floating leaves, as Trapa, Nelumbo, Nymphaea and Marsilea are shown in Figure 4.3. 3, Submerged Floating Hydrophytes They remain in contact with only water, being completely submerged and not rooted in the mud. Their stems are long and leaves generally small. Some of the examples are Ceratophyllu Utricularia, Najas, etc. In Ceratophyllum, (Fig. 4.4), roots are lacking even in émbryo stage, sometimes leafy branches being modified into ‘rhizoids’. 4. Rooted Submerged, Hydrophytes These are hydrophytes like Hydrilla, Potamogeton, Isoetes and Vallisneria (Fig. 45) that remain —CoTHpIstely submerged in water and rooted in soil. (BC-4)64 submerged leaves. Fig. 44, Submerged floating hydrophyte. In Hydrilla, Potamogeton and Chara, the stem is ovig, bearing small Teaves at the nodes. Hydrilla is a slender weed with fibrous roots. In Isoetes ‘and Vallisneria stem is tuberous (corm-like) with Tong leaves, which are narrow, ribbon shaped. 5, Rooted Emergent Hydrophytes They grow in shallow waters. These are the hydrophilous forms which, although require excess of water, but their shoots (assimilatory organs) are partly or completely exposed to air. The root system is completely under water, fixed in soil. In some, as-Sagittaria and Ranunculus (Fig. 4.6) shoots are partly in water and partly emerging ie, exposed to air, whereas in others such as Scirpus and Cyperus (Fig. 4.6) also known as shoots are completely exposed Potamogeton Water Regimes and Ecological Adptation, 1d the importance of soil aj, ind on the basis of the nature gy requirements, divided play number of classes as (i) hydrophytes Cin wats (in marsh); Xerophiloy,” (on acid soils), Gv) psychrophyt, (v) halophytes (on saline soi) (vi) lithophytes (on rocks), (vii) “psammophyts (on sand and gravel), (viii) chersophyte, (on wastelands), (ix) eremophytes (deserts any steppe), (x) psilophytes (on savannah), (xi) seer, phyllous formations (bush and ores, (xii) coniferous formations (forest); any mesophilous— (xii) mesophytes. if in plants’ life soil and water communities, into hydrophilous (i) _helophytes (iii) oxylophytes (on cold soils), ECOLOGICAL ADAPTATIONS IN HYDROPHYTES Hydrophytes may differ from each other in some aspects, but most of the characteristics, morphological as well as anatomical, are common to all of them. Due to these similarities, they belong to the same ecological group-tte hydrophytes. We shall now deal with all such features of these plants which enable them become adapted to such hydrophytic conditions. female female Ise ee Vallisneria Fit. 48, Rooted submerged hydiphyics‘| PLATE I Wotffia Loma Spirodella Salvinia FREE-FLOATING HYDROPHYTESPLATE I Trapa ‘Nymphaea Marsilea ROOTED HYDROPHYTES WITH FLOATING LEAVES Ceratophyllum Utricularia SUBMERGED FLOATING HYDROPHYTESWater Regimes and Ecological Adaptations 65 mergnt eS Ete leaves Gj Ranunculus | — submerged dissected Cyperus Fig. 46. Rooted emergent hydrophytes, Morphological Features Roots Due to availability of water in plenty roots, the principal organs of water absorption, in such plants become of secondary importance, and less significant Their overall development is usually very poor and insignificant in most of the hydrophytes, (1) Roots may be entirely absent as in Wolffia, Salvinia (Fig. 4.2) and Ceratophyllum (Fig. 4.4) or poorly developed as in Hydrilla (Fig. 4.5). In Salvinia (Fig. 4,2) submerged eaves compensate for roots. However, in emergent forms, as Ranunculus (Fig. 4.6) which grow in mud, roots are well developed with distinct root caps. (2) Root hairs are absent or poorly developed. (3) Root caps are usually absent. In some cases, 8 Eichhornia (Fig. 4.2), root caps are replaced by ‘toot pockets’. (4) Roots, if present, are generally fibrous, adventitious, reduced in length, unbranched or poorly branched. In Lemna (Fig. 4.2) roots act simply as balancing and anchoring organs. (5) In some plants, as Jussiaea repens, there develop some floating aerial roots also in addition to normal adventitious roots. Stems (1) In submerged forms as Hydrilla and Potamogeton (Fig. 4.5), stem is long, slender, spongy and flexible, In free-floating forms it may be slender, floating horizontally on water surface as in Azolla (Fig. 4.2) or thick, short, stoloniferous and spongy as in Bichhornia (Fig. 4.2). In forms which are rooted with floating leaves it is a rhizome, as in Nymphaea and Nelumbo (Fig. 43). (2) Vegetative propagation by runners, stolons, stem and root tubers, dormant apices, offsets ec, is most extensive and common method of reproduction, Thus most of them are perennials. (BCA)‘Leaves orms, leavt 1) In submerged form ON er Tong and en ia (Vig. 4.5), oF long Vallisneria (Fig. 4.5 n-ne in Potamogeton (Fig. 4.5), ot finely ti shaped! linear as ed ‘as in Ceratophyllum (Fig. 4.4). Foating Teaves are lange, flat and entire as Nymphaea and Nelimbo (Fig, 4.3) with their a xd with wax; their petioles upper surfaces coate I a jong, flexible, and often covered with imilge In Fiori (Pig. 4.2, and Tropa (Fig. 43) petioles become swollen ani spongy Emergent forms as Ranunculus and Sagittaria Gig. 4.6) show heterophylly with submerged, floating, and acrial leaves. (3) Submerged leaves are generally translucent. Flowers and seeds. These are less common in submerged forms. Where flowers develop, seeds are rarely formed. Anatomical Features Roots The distribution of various tissues in roots of hydrophytes, in general, becomes clear from Figures 4.7 and 4.8 showing transverse sections of roots of Potamogeton and Eichhornia respectively. It may be concluded from these figures that (J) Cuticle is either completely absent or, if present it is thin and poorly developed. Epidermis is usually single-layered made up of thin-walled parenchymatous cells, Cortex is well developed, thin-walled and parenchymatous, major portion of which is occupied by well developed prominent air cavities - the ‘aerenchyma’ which offers resistance to bending stress, incre: buoyancy and allows a rapid gaseous exchange, Vascular tissues are poorly developed least differentiated in submerged forms, as Potamogeton (Fig. 4.7) with thin-welled elements. In xylem, vessels are less common, tracheids being generally present. In floating (BC-4) (2) (3) e and lack (5) Stems Distribution of various tissues in stems hydrophytes becomes clear from transverse sectic? of stem of Hydrilla (Fig, 4.9). It may be se that: wo (2) Fig, 47. TS. root of Potamogeton pectinatus (submerzt hhydrophyte). Note the absence of root hairs and cutci. undifferentiated broad cortex with air chambers, vascalz tissues poorly developed, represented mainly by piloex: of mechanical tissues. forms as Ejichhornia (Fig. 4.8) they comparatively differentiated to some exec However, in emergent forms as Ranun: and Typha etc., vascular elements comparatively much distinct and developed. Mechanical tissues are generally absent exces in some emergent forms, as Typha where pi | cells are sclerenchymatous. Cuticle is either absent or poorly develope! and thin, Epidermis is usually single-layered made 0 of thin-walled parenchymatous _ cell However, rhizomes of Nymphaea 2°water Regimes and Ecological Adaptations 67 middle aerenchyma, S epidermis A ‘outer cortex inner cortex endodermis pericycle xylem phloem pith Fig, 48. TS. root of Eichhornia (floating-leaved hydrophyie). Note, the absence of root hairs and cuticle; undifferentiated well developed cortex with abundance of aerenchyma; mechanical tissues represented only by xylem elements, Nelumbo show well-developed epidermis. In emergent forms as Typha, cuticle as well as ‘a, ‘Pidermis are generally well developed. @) Hypodermis is completely absent in- | Submerged forms as-—-Hydrilla and Potamogeton, However, in floating and ‘mergent forms, it may be present as thin- Walled parenchyma or collenchyma Cortex is well developed, thin-walled and Parenchymatous, extensively traversed by air Cavities as in roots. Cells of cortex generally | Possess chloroplasts and are thus — (5) © @ photosynthetic. In some, as Nymphaea, there are found large number of vascular bundles scattered in the cortex. Endodermis is generally distinct, especially in rhizomes and similar organs. Vascular bundles generally lack bundle sheaths. Vascular elements are thin-walled, lignified elements being absent. However, in , emergent forms as Typha, vascular elements are compa-ratively well differentiated and developed. Mechanical tissues are usually absent.68 Leaves In their internal structure, Ieaves in different variations. However, some of © commion (0 most of in ear from hydrophytes show the anatomical features the eaves. Distrituation of different of hydrophytes would become cl leav Figures 4.10, 4.11 and 4.12 showing vertical transverse sections of leaves of Anacharis, Poramogeton and Nymphaea respectively. From these we may conclude that in leaves: (1) Caticle is usually absent in submerged forms as Anacharis and Potamogeton (Figs. 4.10, 4.11). In floating forms as Nomphaea (Fig. 4.12) it is poorly developed confined only to upper side, and is thin. In emergent forms also it is thin. (2) Epidermis is single-layered, made up of thin- walled cells with abundance of chloroplasts. (3) Stomata are completely absent in submerged Teves as Anacharis and Potamogeton epidermis *hypodermis ericycle Fig, 49. TS, stem of Hydrila (submerged ya wpiifleniid cnen wih ar chanbers, abtnce fs ioe cen somes of trace te xylem being represented only by a cavity in the centre, Water Regimes and Ecological Adaptatio, upper epidermis, parenchyma vascular bundle lower epidermis, mere NOS) Fig, 410, TS, leaf of Anacharis (submerged). Note, i) absence of cuticle and stomata; undifferentiated mesophyy| represented by single-layered parenchyma; reduced vascily tissues represented by thin-walled cells in the cen absence of mechanical tissues. —_ (Figs. 4.10, 4.11). In floating forms Nymphaea (Fig, 4.12) stomata are contin only to the upper surface, of leaf, whereas ix emergent forms they are generally found c both surfaces of leaves. (4) Mesophyll is undifferentiated in submerge: leaves, where it is generally single-layer as in Potamogeton (Fig. 4.11). In Anachar Fig, 4.10) also it is thin and undifferentiated In floating leaves as Nymphaea (Fig. 4.10, it is, however, differentiated into palisade an spongy parenchyma with well-developed sic cavities. In emergent leaves, mesophyll is wel differentiated with air cavities. (5) Vascular tissues are very much reduced and sometimes difficult to be differentiated ian xylem and phloem as in leaves of submerged forms like Anacharis (Fig. 4.10). Whereve| Gifferentiated as in Nymphaea (Fig. 4.12.| xylem elements are thin-walled, phloem beit well developed. However, in’ aerial leaves vascular elements are comparatively vel differentiated with vessels in xylem. upper epidermis feessara ical lower epidermis Teat (only lateral wing port 4 Be 41 ‘wing partion shows) ranoscion pilus (eboerged), Now, the abs | stomata, undifferentiated | ‘mesophyll between two epidermal layers,es and Ecological Adaptations 69 ater Regi cuticle ‘upper epidermis, palisade parenchyma, air chambor tacinay 2 tichosctoreids es “Slower epidermis pprotoxylem lacuna | vascular metaxylem bundle phioom | of id Hb Fig. 412. TS. leaf of Nymphaea (Moating-leaved). Note, the thin cuticle; stomata being confined only to the upper surface; ‘hinewalled epidermal cells; abundance of air chambers in spongy parenchyma; absence of mechanical tissues (only selereids preset reduced vascular elements represented mainly by phloem, xylem being much reduced hair epidermis Jeaved), Note, the absence of cuticle; thin-walled epidermal 1s of collechymatous hypodemmis; abundance ented by Ineuae. Mig. 413. TSS. petiole of Nymphaea (Moating-! ‘ells; reduced mechanical tissues represented only by @ few layer c of aerenchyma; vascular elements with abundance of phloem, xylem being repress N70 (6) Mechanical tissues are abse (7) The petioles, wherever well developed, as it Nymphaea (Fig. 4.13) also possess internally the various tissues characteristic of a typical hydrophyte, ie, an ince of aerenchym, thin-walled cells, lack of differentiation in vascular tissues and absence of any lignified mechanical tissues. MESOPHYTES They are very extensive on the land surface. They are such land plants which grow in moist habitats and need well-aerated soils. They prefer soil and air of moderate humidity and avoid soil with standing water or containing a great abundance of salts. In some respects they stand in between the hydrophytes and xerophytes. Broad-leaved trees growing in wet depressions, along lakes and rivers, are mesophytes. They generally lack the special structural and physiological adaptations found in hydrophytes and xerophytes. The general morpho-anatomical features of mesophyte are as follows: (1) Root system is well developed. Roots are generally fairly branched, with root caps and root hairs. (2) Stems are generally aerial, solid and freely branched. (3) Leaves are generally Jarge, broad, thin and varied in shapes; generally oriented horizontally, green, without hair or waxy coatings (4) Cuticle in all aerial parts moderately developed, (5) Epidermis well developed, without any hair or waxy coatings and cells without chloroplasts. (6) Stomata generally present on both surfaces of leaves. Guard cells show frequent movernents, (7) Mesophyll in leaves is differentiated into Palisade and spongy parenchyma, with many intercellular spaces, P Water Regimes and Ecological Adaptay, (8) Vaseular and sues developed and well differentiated, ne (9) ‘they may show temporary wilting gy, noon hours. i XEROPHYTES | ‘The term xerophyte has been defined ,, interpreted variously. Morphological features rates of transpiration have almost failed explain the truc nature of this group of pla ‘They are sometimes loosely defined as ‘plant; » dry habitats’. But if taken in this sense, mos the mesophytes should be called xerophy Daubenmire (1959) in his discussion on yi, group defined xerophytes as ‘plants which gro on substrata that usually become depleted growth water to a depth of at least 2 decimes during a normal season’. Thus, in arid zones, plants not confined to the margins of streams lakes have been considered as xerophyt whereas in regions of heavy rainfall the cl would be represented only by shallow-roozi plants of sandy soils, by plants of dry ridgetop. and by algae, mosses and lichens which gov on tree barks or rock surfaces etc. The nature of xerophytism is not clearly understooé For example, it is difficult to decide whether 2 xerophyte is really xerophilous and occurs oz} in dry habitats and deserts, or is merely drough resistant. There are instances, for example, tt if desert plants are grown in soils with moder? amount of water, they show better growth as Artemisia fasciculata, Zygophyllum fabago Centaurea solstitialis, the yield of _ plan increased to 1,5-2 times by increasing wate holding capacity of soil from 40% to 60% Thus, such xerophytes are not xerophilous # least so far as the soil moisture is concemel Oppenheimer (1960), in a literature survel favours a much broader interpretation includité all the — morphological, anatomical Physiological modifications which may assist plant to cope with environmental water deficitwater Regimes and Ecological Adaptations ‘On the basis of their morphology, physiology id life cycle pattern, xerophytes are generally Zpssified into the following (hr 1. Ephem “drought evaders’ or drought escapers’. They are mostly found in arid ones. They are aanvals, which complete their fife Gyeles within a very short period of 6-8 weeks or Se. With their small size and large shoots in felation 10 roots, they are well adapted 10 such dry habitats. They actually avoid and not withstand dry seasons, and thus escape dryness in external and infernal environments. Some do not prefer {o call them true xerophytes. Examples—Argemone mexicana, Solanum xanthocarpum, Cassia tora, ‘Tephrosia purpurea, etc. 2. Succulents ‘They are the plants that suffer from dryness in external environment only. Their succulent, fleshy organs (stems, leaves, roots) serve as water-storage organs which accumulate large amount of water n system is shallow, stem swollen and leaves thick, leathery and succulent. Some of the examples are Aloe, Euphorbia and Opuntia (Fig. 4.14) and Various cacti, Agave and Ceiba parviflora, AS the succulents avoid drought, some prefer to exclude them from true xetophytes. This is indeed a unique mode of adaptation, In some of them, stems become succulent, which are also called the ‘fleshy xerophytes’ as in Opuntia and Euphorbia (Fig. 4.14). In such xerophytes, cuticle is thick, and well developed hypodermis is two-to-three layered, collenchy- matous; cortex is also thick-walled with chlorenchymatous cells, below which there is present a prominent ‘water-storage region’, whose cells are thin-walled with a few intercellular spaces. Those succulent xerophytes in which leaves become fleshy; are also known as ‘malacophyllous xerophytes’, such as Aloe (Fig. 4.14), Begonia, Salsola, Bryophyllum, Agave; Yucca, Tradescantia etc. Their leaves possess turgescent parenchy- matous cells with thin walls. Asparagus and Ceiba parvifolia store water in their roots. Euphorbla ‘Succulent _xerophytes X spines a 4 hylloctadeWater Regimes and Ecological Adaptation, = Uitfer in soveral ways from each other bye 5 my all possess cortain featur 3. Non-Succulent Perenn my iy " Mt common, On the basis of thes f sx a number of grouped into the same ecological group ~ 4 dear nd physiological —erophytes. These features, which are more or | phys to all, and which enable them conditions, a are actually the trie xerophytes or droup. Ms, beet ‘morphological natomieal andl em to withstand common w characteristics which enable them (0 wi yal, a whic Stat iy conto Tey. the plans that guyive une the preveling airy. conditions 4 : " + from dryness hott in their infernal as well as follows: is. ‘Their morpho- ; Morph 1 Features lo; i ester en he ji i re and extensive root system, high osmotic pr enturance of desiccation, ability (0 reduce Rggtg extremely level during transpiration to an extremely. low f : permanent wilting, and reduction in size of leaf fn contrast with hydrophytes which develop jy blades and cells, Non lent perennials like conditions with plenty of water, xerophytes develop Caloiropis procera, Acacia nelotica, Zizyphus under water-deficient conditions. Thus, in order jy jgitha and Capparis apyla are shown in Figute secure water, which is present in less amount an, 4.15. Other examples are Prosopis, Casuarina, moreover, in deeper lavers of soil, roots in Nerium, Alhagi, Saccharum, and Salvadora. xerophytes become the principal organs of primary ‘The general characteristic features of non jmportance. The root system is thus very we) succulent xerophytes are as follows: developed, with the following characteristics: (1) Root system is very extensive. For example, (1) Ieis very extensive, which in some cases i in Prosopis, Calotropis and Alfalfa, roots may several times longer than the shoot. Roos be more than 30 meters Jong, are long, tap roots, with extensive branching (2) Following characteristics help minimizing the spread over wide areas, rate of transpiration: (i) dying back of leaves @ as in many grasses; (i) rolling and folding of leaves as in many grasses like Ammophila arenaria; (ii) delicate leaves which are shed under conditions of less supply of ‘water: Ge) heavy cuticular and epidermal layers: £2) waxy coatings on Teaves; sunken type a vey they are stunted, Woody, dry, hard, Of stomata on leaves; (vii) very small, natrow, dged and covered with thick bark, sarah Say leaves; (vi) ridged stems, 2) In some as Saccharum, stem vecome Root hairs and root caps are very well developed. Stems (3) Aerial organs may become variously modified underground, whereas in Opuntia (Fig. 4.14) according to the prevailing climatic it becomes fleshy, green, —_leaf-like eens (hylloclade) covered with spines. In *perimentally, the reduction of water supply Euphorbia also (Fig. 4.14) it becomes fleshy fis been shown to reduce the shootroot ratio and green, in some grasses 3) On stems and leaves there are generally hairs = and/or waxy coatings, ECOLOGICAL ADA LeavesPLATE I Hi\\ ROOTED SUBMERGED HYDROPHYTES Sagittaria ROOTED EMERGENT HYDROPHYTESPLATE IV Aloe Euphorbia SUCCULENT XEROPHYTES Zizyphus NON-SUCCULENT XEROPHYTESwater Regimes and Ecological Adaptations 73 qj —Noral buds stipular 4 spines \\ Jf as leaves stipular spines SS Capparis, wuceulent perennials ~ 2) Foliage leaves, wherever present, may become © Leaf surfaces are generally shiny and glazed thick, fleshy ‘and succulent, or tough and to reflect light and heat ' leathery in texture. : (4) In some monocots as Ammophila, Poa, and i ‘Agropyron (Fig. 4.21 A-C), leaves becomeRegimes and Ecological Adaptatio, Water 74 utile c epidermis hypodermis * (parenchymatous) cortical vascular bundle ‘starchy sheath Phloem ] stelar ‘cambium | vascular bundle ‘xylem Fig, 416. T'S. stem (a part) of Cawarina. Note the thick cuticle; sunken stomata, conlined ayn Froawes: Presence of hairs in grooves; sclerenchymatous hypodermis, green palisade region of subhypodermal cortex; well-developed vascular and mechanical tiscues, folded and rolled in such a manner that the sunken stomata become hidden, and thus rate of transpiration is considerably minimised, ©) In some of them as Euphorbia (Fig, 414), Acacia nelotica, Zizyphus jujuba and | Capparis aphylla (Fig. “4.15), Stipules become modified into spines, water 1 ‘storage cells Palisade upper epidermis Anatomical F ‘eatures —— i Matures ‘spongy . lower at the root tips, ©) Roots may becom re a in Asparagus ® Im Pinus edu ee and Calotropi; Possess rigid and ae thickened walls,water Regimes and Ecological Adaptations 75 x substomatal. chamber vascular, bundles SMa aa vascular bundles stoma _ guard coll epidermis —wator storage tissue cuticle proper | ‘cuticle (cutinised) layer | outer wall of ‘epidermal cell -water storage tissue Fig, 418 7S, leaf of Aloe (uceulend. A. T and enlarged. Note, the well developed centrally placed parenchymatous thin-walled cells located in between Stems (1) In succulent xerophytes, stems possess ® Water-storage region. In stems of most of the non-succulent ae such as Casuarina ‘ig. 4.16), there are present the following chief characteristics: snmais Ty Cenral part of the same cell | the upper and lower palisade lay tatile; sunken stomata; thickened outer walls of epider tissue made up of urs water storage escent ick eloped palisae le is very thick well developed, with heavily alls. several-layered « Hypodermis and sclerenchymatous. « Stomata are of sunken (ype. ‘ular tissues are very well developed, heavily lignified. VascularWater Regimes and Ecological Adeptati,, 76 (2) Leaves of non-succulent xerophytes, bundles have well developed several- layered bundle sheaths. ‘s Mechanical tissues are very well developed. 3) Bark is very well developed. (4) Oil and resins are often present. Leaves (1) In succulent Ieaves of malacophyllous xerophytes, such as Peperomia (Fig. 4.17), epidermal cells of leaves serve as water- storage organs. Similarly, succulent leaves of Aloe (Fig. 4.18), and Salsola have prominent water-storage regions in their mesophyll. Moreover, in such leaves (Figs. 4.18, 4.22) cuticle is thick and outer walls of the epidermal cells are heavily deposited with cutin and cellulose, vascular bundle : of vein mesophyll — xylem, bundle spongy palisade sheath Parenchyma parenchyma oxalate cuticle as Nerium (Fig. 4.19) and Pinus (Fig. 4) elc., possess: a Well developed heavy cuticle. # Several-layered epidermis in Neriun, a several-layered, sclerenchymatoy, hypodermis in Pinus. ‘+ Mesophyll very well differentiated iny, palisade and spongy parenchyma, @ Stomata of sunken type confined only 1 lower epidermis. In some xerophytes ay Nerium (Fig. 4.19), stomata are situate in pits lined with hairs. © Vascular tissues very well developed, differentiated into xylem with lignified elements, and phloem. In Neriwm, ip addition to big vascular bundle in mid-ip region, there are several other vascular bundles also. calcium, upper eystals epidermis cuticle lower epidermis vascular bundle of mid ribter Regimes and Ecological Adaptations wat (sclerenchymatous) Be 420, TS. peedle of Pinus roxburghii (con-succulent perennial), Note, thick cule) Wicicwalled epldconi Secmymaious bypodermis; sunken stomata; mesophyll cells with inflding; complex transfusion tissue; well developed vascular tissues with abundance of xylem elements, vascular bundle cuticle sclerenchyma Jower epidermis parenchyma photosynthetic tissue cuticle stomata lower epidermis -sclerenchyma ‘vascular bundle “per epidermis. Horenchyma—_ ef Leaf roling/olding devices in some non-sueculenl perennials Cross sections (agrammaic) of rolled/olded jrnophita arenaria (A) where the laf blade is rolled inwards with ridges and grooves on its adaxial side, thick ba ied to upper epidermis only, which came to lie in a hidden position due to leaf rolling; selerenchymatous SS Arroryon op (3) whew the if rolled iu tbe sama mama asin (A) and nolo the pence of ek ig tissues, hairs, hidden stomata, etc. ze78 ical tissues very well developed, « Mechanii ‘ including several kinds of sclereids. In Pinus, there is well developed complex transfusion tissue. ; In some non-succulent xerophytes, particularly ~ grasses, as Ammophila and Agropyron (Fig. 4.21 A-B), leaves become rolled and/or folded in such a manner that the stomata occupy a hidden position, thus mi ing the rates of transpiration. Moreover, their epidermal cells remain mostly turgid. We have described in detail the various morphological and anatomical features of each of the three major ecological groups of plants, hydrophytes, mesophytes and xerophytes. A comparative account of the same is presented in Table 4.1 also. HALOPHYTES ‘They are a special type of xerophilous plants which grow on saline soils with high concentration of salts like NaCl, MgCly and MgSO,. Soils are physiologically dry. ‘Their osmotic pressure values are very high. Most of them are succulents, their leaves are evergreen, thin, small and leathery with water storage tissues, thick cuticle and well developed palisade Water Regimes and Ecological Adaptatig,, ‘i Most of them produce special type ,- branched, negatively geotropic roots that con, 7 out of the mud surface to encourage the ony of oxygen gas. These roots are cal, pneumatophores which possess breathing po for gascous exchange. Some of the halophyg, show vivipary ie., germination of seeds befo, they are shed from parent plant. The univers: feature of these plants is selective absorption ang tolerance of sodium chloride, and the ability \, grow in soils so saline that they are cith, physiologically toxic to normal plants or impos so serious an osmotic water stress that wate absorption is impeded. Plant cells are bathed jy a medium of high osmotic potential and thus should maintain a higher vacuole ‘Osmotic potential in order to avoid loss of water. Halophytic communities have _ been divided (Warming, 1909) into (i) lithophilous, (i) psammophilous, (iii) pelophilous, ani (iv) helophilous types, growing on rock and stones, sand, mud, and swamp respectively Helophilous halophytes have been further sub. divided into (i) salt-swamp and salt-desert, and Gi) littoral swamp-forest (mangrove) types, of which the latter are most extensive, which occur in all tropical seas, especially on flat, muddy shores, where the water is relatively calm, as in lagoons, inlets, estuaries etc, Soil is flooded with water either permanently or at high tide. These epistomatal chamber stoma Cuticle proper outer wall of cutinised layer epidermal cell cellulose layerwater Regimes and Ecological Adaptations pate ydrophtes ee enc ms Bet cc ~ ‘podified into rhizomes, thin, delicate, wie orn noe ee ie oom ttl een ant al Mesophytes + Root system wel developed, tp or brows roots with root hari, Stem is rigid and stout, 2. Leaves large, thin, gen Kare Imps, e,generty wont 3. Leaves sel Acrenchyma lacking. 1 79 Xerophytes Root sytem” smelly as cs oig ret tal sre! tine np the ht hard, woody, . : mich dae, tometer nderpond branched, ‘sometimes much rediueed to scales or modified into spines, in some leathery and thick with shining surface and waxy ‘couting, covered with hairs Aerenchyma lacking. vegetative organs. erative OF 2. Cuticle developed. 2. Cuticle thick, well developed. 2, Cuticle generally absent 3. Stomata on one or both the surfaces. 3, = stomata either absent, or if present, ‘nly at upper surface or even non- funetonal. ‘4. Mesophyil undifferentiated into 4, Palisade |. Stomata less in number, generally confined to lower surfaces of leaves, sunken. ‘well developed and 4. Palisade generally on both sides of tals and spongy parenchyma. differentiated. : i leaves, ces and vacuoles small. 5, Colorophyl in addition to leaves also 5. Chlorophyll mostly in leaves m in other parts of plant. pl mostly in leaves 5. Gop! mowty ie sem et 6 Epidermal calls thin-walled. 6. Epidermal cells thick ide eon a ae 6, Epidermal es omcssy ik conducting elements very few, nom 7. Meachanical and vascular tissues 7, Mechanical and vascular tssues very lignfied. well developed. Physiological Habitats mostly deficient in oxygen content, thus having an ability to temporary respre anaerobically, or have low temperature. ‘oxygen requirements. Special seraling organs are present. litoral swamp-forests form a characteristic Vegetation, known as the mangroves or mangrove formation in tropical and subtropical areas. In India, mangroves are very much prevalent n seashores of Bombay and Kerala and in os and Nicobar islands. They grow on water- : ea soil, where habitat is characterised by sandy. anny and saline soils, high precipitation, high i spheric humidity and almost no fluctuation 1" apenature throughout the year. The wo common Phytes of this type are Rhizophora mucronata ean eratia. They produce stilt roots, reno phores, and show vivipary. They show wrphic characters as evident from the ‘Normal physiological processes, with wilting at room well developed. Most of the features directed 10 reduce the anspiration rates: cephemerals complete life cycle ia shore period, High osmotic pressure (as much as 3100 atms. in some halophytes), and endurance of desiccation, anatomical characters of vegetative organs of Rhizophora mucronata given below. Roots Roots are of two types, aerial and subterranean. ‘The subterranean root in transverse section (Fig. 4.23) shows: (1). Several-layered cork. @) Comtex mnde up of | starshaped oF stellate cells, connected with each other by Tateral arms, Cortical cells have peculiar thickening ridges which are lignified. Some tells of cortex are filled with oil and and tannin. (BC-4)